This general sense gave way to the modern English sense by the end of the Middle English period, around 1500. However, all modern Germanic languages save English and Scots retain the more general sense: for example, German Tier, AlemannicDiere or Tiere, Pennsylvania DutchGedier, Dutch dier, Afrikaansdier, Limburgishdiere, Norwegian dyr, Swedish djur, Danish dyr, Icelandic dýr, Faroesedýr, West Frisiandier, and North Frisiandiarten, all of which mean "animal". (However, contrary to south European languages, Dama in Latin and daim in French mean "fallow deer" only).[2]

For most types of deer in modern English usage, the male is called a "buck" and the female is termed a "doe", but the terms vary with dialect, and especially according to the size of the species. For many larger deer, the male is termed a "stag", while for other larger deer the same words are used as for cattle: "bull" and "cow". The male red deer is a "hart", especially if more than five years old, and the female is a "hind", especially if three or more years old; both terms can also be used for any species of deer, and were widely so used in the past.[3] Terms for young deer vary similarly, with that of most smaller species being called a "fawn" and that of most larger species "calf"; young of the smallest kinds may be a kid. A castrated male deer is a "havier". A group of deer of any kind is a "herd". The adjective of relation pertaining to deer is cervine; like the family name "Cervidae", this is from Latin: cervus, "deer".[4]

Habitat

Deer live in a variety of biomes, ranging from tundra to the tropical rainforest. While often associated with forests, many deer are ecotone species that live in transitional areas between forests and thickets (for cover) and prairie and savanna (open space). The majority of large deer species inhabit temperate mixed deciduous forest, mountain mixed coniferous forest, tropical seasonal/dry forest, and savanna habitats around the world. Clearing open areas within forests to some extent may actually benefit deer populations by exposing the understory and allowing the types of grasses, weeds, and herbs to grow that deer like to eat. Additionally, access to adjacent croplands may also benefit deer. However, adequate forest or brush cover must still be provided for populations to grow and thrive.

Deer are widely distributed, with indigenous representatives in all continents except Antarctica and Australia, though Africa has only one native deer, the Barbary stag, a subspecies of red deer that is confined to the Atlas Mountains in the northwest of the continent. However, fallow deer have been introduced to South Africa. Small species of brocket deer and pudús of Central and South America, and muntjacs of Asia generally occupy dense forests and are less often seen in open spaces, with the possible exception of the Indian muntjac. There are also several species of deer that are highly specialized, and live almost exclusively in mountains, grasslands, swamps, and "wet" savannas, or riparian corridors surrounded by deserts. Some deer have a circumpolar distribution in both North America and Eurasia. Examples include the caribou that live in Arctic tundra and taiga (boreal forests) and moose that inhabit taiga and adjacent areas. Huemul deer (taruca and Chilean huemul) of South America's Andes fill the ecological niches of the ibex and wild goat, with the fawns behaving more like goat kids.

The highest concentration of large deer species in temperate North America lies in the Canadian Rocky Mountain and Columbia Mountain regions between Alberta and British Columbia where all five North American deer species (white-tailed deer, mule deer, caribou, elk, and moose) can be found. This region has several clusters of national parks including Mount Revelstoke National Park, Glacier National Park (Canada), Yoho National Park, and Kootenay National Park on the British Columbia side, and Banff National Park, Jasper National Park, and Glacier National Park (U.S.) on the Alberta and Montana sides. Mountain slope habitats vary from moist coniferous/mixed forested habitats to dry subalpine/pine forests with alpine meadows higher up. The foothills and river valleys between the mountain ranges provide a mosaic of cropland and deciduous parklands. The rare woodland caribou have the most restricted range living at higher altitudes in the subalpine meadows and alpine tundra areas of some of the mountain ranges. Elk and mule deer both migrate between the alpine meadows and lower coniferous forests and tend to be most common in this region. Elk also inhabit river valley bottomlands, which they share with White-tailed deer. The White-tailed deer have recently expanded their range within the foothills and river valley bottoms of the Canadian Rockies owing to conversion of land to cropland and the clearing of coniferous forests allowing more deciduous vegetation to grow up the mountain slopes. They also live in the aspen parklands north of Calgary and Edmonton, where they share habitat with the moose. The adjacent Great Plains grassland habitats are left to herds of elk, American bison, and pronghorn antelope.

The Eurasian Continent (including the Indian Subcontinent) boasts the most species of deer in the world, with most species being found in Asia. Europe, in comparison, has lower diversity in plant and animal species. However, many national parks and protected reserves in Europe do have populations of red deer, roe deer, and fallow deer. These species have long been associated with the continent of Europe, but also inhabit Asia Minor, the Caucasus Mountains, and Northwestern Iran. "European" fallow deer historically lived over much of Europe during the Ice Ages, but afterwards became restricted primarily to the Anatolian Peninsula, in present-day Turkey.

Present-day fallow deer populations in Europe are a result of historic man-made introductions of this species, first to the Mediterranean regions of Europe, then eventually to the rest of Europe. They were initially park animals that later escaped and reestablished themselves in the wild. Historically, Europe's deer species shared their deciduous forest habitat with other herbivores, such as the extinct tarpan (forest horse), extinct aurochs (forest ox), and the endangered wisent (European bison). Good places to see deer in Europe include the Scottish Highlands, the Austrian Alps, the wetlands between Austria, Hungary, and the Czech Republic and some fine National Parks, including Doñana National Park in Spain, the Veluwe in the Netherlands, the Ardennes in Belgium, and Białowieża National Park of Poland. Spain, Eastern Europe, and the Caucasus Mountains still have virgin forest areas that are not only home to sizable deer populations but also for other animals that were once abundant such as the wisent, Eurasian lynx, Iberian lynx, wolves, and brown bears.

The highest concentration of large deer species in temperate Asia occurs in the mixed deciduous forests, mountain coniferous forests, and taiga bordering North Korea, Manchuria (Northeastern China), and the Ussuri Region (Russia). These are among some of the richest deciduous and coniferous forests in the world where one can find Siberian roe deer, sika deer, elk, and moose. Asian caribou occupy the northern fringes of this region along the Sino-Russian border.

The highest concentration of large deer species in the tropics occurs in Southern Asia in Northern India's Indo-Gangetic Plain Region and Nepal's Terai Region. These fertile plains consist of tropical seasonal moist deciduous, dry deciduous forests, and both dry and wet savannas that are home to chital, hog deer, barasingha, Indian sambar, and Indian muntjac. Grazing species such as the endangered barasingha and very common chital are gregarious and live in large herds. Indian sambar can be gregarious but are usually solitary or live in smaller herds. Hog deer are solitary and have lower densities than Indian muntjac. Deer can be seen in several national parks in India, Nepal, and Sri Lanka of which Kanha National Park, Dudhwa National Park, and Chitwan National Park are most famous. Sri Lanka's Wilpattu National Park and Yala National Park have large herds of Indian sambar and chital. The Indian sambar are more gregarious in Sri Lanka than other parts of their range and tend to form larger herds than elsewhere.

The Chao Praya River Valley of Thailand was once primarily tropical seasonal moist deciduous forest and wet savanna that hosted populations of hog deer, the now-extinct Schomburgk's deer, Eld's deer, Indian sambar, and Indian muntjac. Both the hog deer and Eld's deer are rare, whereas Indian sambar and Indian muntjac thrive in protected national parks, such as Khao Yai. Many of these South Asian and Southeast Asian deer species also share their habitat with other herbivores, such as Asian elephants, the various Asian rhinoceros species, various antelope species (such as nilgai, four-horned antelope, blackbuck, and Indian gazelle in India), and wild oxen (such as wild Asian water buffalo, gaur, banteng, and kouprey). One way that different herbivores can survive together in a given area is for each species to have different food preferences, although there may be some overlap.

Australia has six introduced species of deer that have established sustainable wild populations from acclimatisation society releases in the 19th century. These are the fallow deer, red deer, sambar, hog deer, rusa, and chital. Red deer introduced into New Zealand in 1851 from English and Scottish stock were domesticated in deer farms by the late 1960s and are common farm animals there now. Seven other species of deer were introduced into New Zealand but none are as widespread as red deer.[5]

Description

Deer weights generally range from 30 to 300 kg (70 to 700 lb). The smallest species, the northern pudú, averages 10 kg (20 lb) and the largest, the moose, averages 431 kg (1,000 lb). They generally have lithe, compact bodies and long, powerful legs suited for rugged woodland terrain. Deer are also excellent jumpers and swimmers. Deer are ruminants, or cud-chewers, and have a four-chambered stomach. The teeth of deer are adapted to feeding on vegetation, and like other ruminants, they lack upper incisors, instead having a tough pad at the front of their upper jaw. Some deer, such as those on the island of Rùm,[6] do consume meat when it is available.[7]

The Chinese water deer, tufted deer, and muntjac have enlarged upper canine teeth forming sharp tusks, while other species often lack upper canines altogether. The cheek teeth of deer have crescent ridges of enamel, which enable them to grind a wide variety of vegetation.[8] The dental formula for deer is: 0.0-1.3.33.1.3.3

Nearly all deer have a facial gland in front of each eye. The gland contains a strongly scented pheromone, used to mark its home range. Bucks of a wide range of species open these glands wide when angry or excited. All deer have a liver without a gallbladder. Deer also have a tapetum lucidum, which gives them sufficiently good night vision.

Antlers

With the exception of the Chinese water deer, which have tusks, all male deer have antlers. Sometimes a female has a small stub. The only female deer with antlers are reindeer (caribou). Antlers grow as highly vascular spongy tissue covered in a skin called velvet. Before the beginning of a species' mating season, the antlers calcify under the velvet and become hard bone. The deer rubs off the velvet, leaving dead bone which forms the hard antlers. After the mating season, the pedicle and the antler base are separated by a layer of softer tissue, and the antler falls off.

One way that many hunters are able to track main paths that the deer travel on is because of their "rubs". The deer rub trees to deposit scent from glands near the eye and forehead and physically mark territory.

During the mating season, bucks use their antlers to fight one another for the opportunity to attract mates in a given herd. The two bucks circle each other, bend back their legs, lower their heads, and charge. The tines on the antlers create grooves that allow another male's antlers to lock into place. This allows the males to wrestle without risking injury to the face.[9]

Antlers can be a sign of genetic quality. Males with larger antlers relative to body size tend to have increased resistance to pathogens [10] and higher reproductive capacity.[11] Necropsy research on wild deer that were killed and eaten by wolves shows that deer with asymmetric antlers are weakened by genetic defects and are less likely to escape being caught by predators.[citation needed]

Each species has its own characteristic antler structure – for example white-tailed deer antlers include a series of tines sprouting upward from a forward-curving main beam, while fallow deer and moose antlers are palmate, with a broad central portion. Mule deer and black-tailed deer, species within the same genus as the white-tailed deer, have bifurcated (or branched) antlers—that is, the main beam splits into two, each of which may split into two more.[12] Young males of many deer, and the adults of some species, such as brocket deer and pudus, have antlers that are single spikes.

Cervocerus novorossiae

Colour

Piebald deer

Piebald fawn

Piebald doe

A piebald deer is a deer with a brown and white spotting pattern that is not caused by parasites or diseases. They can appear to be almost entirely white. In addition to the non-standard coloration, other differences have been observed: bowing or Roman nose, overly arched spine (scoliosis), long tails, short legs, and underbites.

White deer

Seneca County, New York maintains the largest herd of white deer. White pigmented white-tailed deer began populating the deer population in what is now known as the Conservation Area of the former Seneca Army Depot. The U.S. Army gave the white deer protection while managing the normal-colored deer through hunting. The white deer coloration is the result of a recessive gene.

White tail fawns are born a brown or tan color with a spotted white pattern. Sometimes these fawns can be born with a grey appearance, making them seem dirty. The coats become pure white in the middle of their second year, and they are sometimes mistaken for albino deer.

Albino whitetail deer appear to have pink skin with a pure white coat, and the irises of their eyes are usually pink as well. There is no such thing as a partial albino; true albino deer have little or no melanin in their bodies. Their color is mainly white because it lacks any pigments, making the skin appear pink because the flowing blood can be seen through the skin. Their white coats make them especially vulnerable to predators. [14]

Biology

Diet

Deer are browsers, and feed primarily on leaves. They have small, unspecialized stomachs by ruminant standards, and high nutrition requirements. Rather than eating and digesting vast quantities of low-grade fibrous food as, for example, sheep and cattle do, deer select easily digestible shoots, young leaves, fresh grasses, soft twigs, fruit, fungi, and lichens. The low-fibered food, after minimal fermentation and shredding, passes rapidly through the alimentary canal. The deer require a large amount of minerals such as calcium and phosphate in order to support antler growth, and this further necessitates a nutrient-rich diet.

Reproduction

Nearly all cervids are so-called uniparental species: the fawns are only cared for by the mother, known as a doe. A doe generally has one or two fawns at a time (triplets, while not unknown, are uncommon). The gestation period is anywhere up to ten months for the European roe deer. Most fawns are born with their fur covered with white spots, though in many species they lose these spots by the end of their first winter. In the first twenty minutes of a fawn's life, the fawn begins to take its first steps. Its mother licks it clean until it is almost free of scent, so predators will not find it. Its mother leaves often to graze, and the fawn does not like to be left behind. Sometimes its mother must gently push it down with her foot.[15] The fawn stays hidden in the grass for one week until it is strong enough to walk with its mother. The fawn and its mother stay together for about one year. A male usually leaves and never sees his mother again, but females sometimes come back with their own fawns and form small herds.

Evolution

The evolution of deer took about 30 million years. There are not many prominent fossils to trace this evolution, but only fragments of skeletons and antlers that might be easily confused with false antlers of non-cervid species.[16] Biologist Valerius Geist suggests evolution to have taken place in stages. The earliest stage featured a poorly defined omnivore of the Palaeogene, that gave rise to deer-like species that lacked antlers but bore horns, such as Protoceras, towards the Neogene. The Protoceras was 1 m (3.3 ft) tall, with small horns. Syndyoceras also shared similar features common with the deer, horse, giraffe, and antelope. Fossils dated approximately 35 million years ago, which were found in North America, show it had bony skull outgrowths that resembled non-deciduous antlers.[17]

By the late Oligocene, saber-toothed deer had evolved from gelocids (early pecoran ruminants). In the early Miocene, muntjacs, the first antler-bearing deer, made their appearance in Eurasia. In 1994, fossils of a large muntjac, Megamuntjacus vuquangensis, were excavated in the Annamite Range(Vietnam). Similar evidence has also been discovered in Laos and Cambodia. The early muntjacs varied in size - as small as hares or as large as fallow deer. They had tusks for fighting and antlers for defense.[16]

Later species were often larger, with more impressive antlers, and, in many cases, no upper canine teeth. They rapidly spread to the other continents. For a time they occupied much of northern Africa, but are now almost wholly absent from that continent. Some extinct deer had huge antlers, larger than those of any living species. Examples include Eucladoceros, and the giant deer Megaloceros, whose antlers stretched to 3.5 m (11 ft) across. Another animal also thought to be related to the deer is the world's oldest known antler-shedding deer, Dicrocerus elegans. This animal's sediment deposits are found in European soil dating back to between 15–30 million years ago.[18] Yet another distant ancestor is thought to be Archaeomeryx.

Taxonomy

Till the end of the twentieth century, it was assumed that the family Moschidae (musk deer) was a sister group to Cervidae. However, molecular analysis has revealed that Moschidae has a closer relationship to Bovidae rather than to Cervidae. It has been estimated that Cervidae diverged from Moschidae and Bovidae 27 to 28 million years ago.[19]

Extant subfamilies, genera and species

The deer family has at least 90 species; the list is based on several molecular and phylogenetic studies by Elisabeth Vrba, Colin Groves and Peter Grubb among others. Note that the terms indicate the origin of the groups, not their modern distribution: the water deer, for example, is a New World species but today is found only in China and Korea. The family Cervidae is organized as follows:[20][21][22][23][24][25]

Hybrid deer

In On the Origin of Species (1859), Charles Darwin wrote "Although I do not know of any thoroughly well-authenticated cases of perfectly fertile hybrid animals, I have some reason to believe that the hybrids from Cervulus vaginalis and Reevesii [...] are perfectly fertile." These two varieties of muntjac are currently considered the same species.

A number of deer hybrids are bred to improve meat yield in farmed deer. American elk (or wapiti) and Red Deer from the Old World can produce fertile offspring in captivity, and were once considered one species. Hybrid offspring, however, must be able to escape and defend themselves against predators, and these hybrid offspring are unable to do so in the wild state. Recent DNA, animal behavior studies, and morphology and antler characteristics have shown there are not one but three species of Red Deer: European red deer, Central Asian red deer, and American elk. The European elk is a different species and is known as moose in North America. The hybrids are about 30% more efficient in producing antlers by comparing velvet to body weight. Wapiti have been introduced into some European red deer herds to improve the Red Deer type, but not always with the intended improvement.

In New Zealand, where deer are introduced species, there are hybrid zones between red deer and North American wapiti populations and also between red deer and sika deer populations. In New Zealand, red deer have been artificially hybridized with Père David deer in order to create a farmed deer that gives birth in spring. The initial hybrids were created by artificial insemination and back-crossed to red deer. However, such hybrid offspring can only survive in captivity free of predators.

In Canada, the farming of European red deer and red deer hybrids is considered a threat to native wapiti. In Britain, the introduced sika deer is considered a threat to native red deer. Initial sika deer/red deer hybrids occur when young sika stags expand their range into established red deer areas and have no sika hinds to mate with. They mate instead with young red hinds and produce fertile hybrids. These hybrids mate with either sika or red deer (depending which species is prevalent in the area), resulting in mongrelization. Many of the sika deer that escaped from British parks were probably already hybrids for this reason. These hybrids do not properly inherit survival strategies and can only survive in either a captive state or when there are no predators.

In captivity, mule deer have been mated to white-tail deer. Both male mule deer/female white-tailed deer and male white-tailed deer/female mule deer matings have produced hybrids. Less than 50% of the hybrid fawns survived their first few months. Hybrids have been reported in the wild but are disadvantaged because they don't properly inherit survival strategies. Mule deer move with bounding leaps (all four hooves hit the ground at once, also called "stotting") to escape predators. Stotting is so specialized that only 100% genetically pure mule deer seem able to do it. In captive hybrids, even a one-eighth white-tail/seven-eighths mule deer hybrid has an erratic escape behaviour and would be unlikely to survive to breeding age.[citation needed] Hybrids do survive on game ranches where both species are kept and where predators are controlled by man.

In history

After the evolution of the hominids, deer became the primary source of food for them. In China, the Homo erectus fed upon the sika deer, the red deer was hunted in Germany. In the Upper Palaeolithic, the reindeer was the staple food for the Cro-Magnon people. Deer had a central role in the ancient art, culture and mythology of the Hittites, the Ancient Egyptians, the Celts, the ancient Greeks, the Asians and several others. In Japanese Shintoism, the sika deer is believed to be a messenger to the gods. In China deer are associated with great medicinal significance, while spotted deer in particular are believed to accompany the god of longevity. Deer was the principal sacrificial animal for the Huichal Indians of Mexico. In medieval Europe, deer appeared in hunting scenes and coats-of-arms. Deer are depicted in many materials by various pre-Hispanic civilizations in the Andes.[28][29]

In literature

Deer have been an integral part of fables and other literary works since the inception of writing. Stags were used as symbols in the latter Sumerian writings. For instance, the boat of Sumerian god Enki is named the Stag of Azbu. There are several mentions of the animal in the Rigveda as well as the Bible. In Indian epic Ramayana, Sita is lured by a golden deer which Rama tries to catch. In absence of both Rama and Lakshman, Ravana kidnaps Sita. In many of his allegorical fables such as "The Stag at the Pool", "The One-Eyed Doe" and "The Stag and a Lion", the legendary Greek author Aesop personifies deer to give moral lessons. For instance, "The Sick Stag" gives the message that uncaring friends can do more harm than good.[28] The Yaqui deer song accompanies the deer dance which is performed by a pascola [from the Spanish 'pascua', Easter] dancer (also known as a deer dancer). Pascolas would perform at religious and social functions many times of the year, especially during Lent and Easter.[28][30]

Economic significance

Deer have long had economic significance to humans. Deer meat, for which they are hunted and farmed, is called venison. Deer organ meat is called humble (see humble pie). The skins make a peculiarly strong, soft leather, known as buckskin. There is nothing special about skins with the fur on since the hair is brittle and soon falls off. The hoofs and horns are used for ornamental purposes, especially the antlers of the roe deer, which are utilized for making umbrella handles, and for similar purposes; elk horn is often employed in making knife handles. In China, a medicine is made from stag horn, and the antlers of certain species are eaten when "in the velvet".[34]

Deer have long been bred in captivity as ornaments for parks, but only in the case of reindeer has thorough domestication succeeded.[34] The Sami of Scandinavia and the Kola Peninsula of Russia and other nomadic peoples of northern Asia use reindeer for food, clothing, and transport. Deer bred for hunting are selected based on the size of the antlers.[35]

The caribou in North America is not domesticated or herded as is the case of reindeer (the same species). Reindeer are often found in colder regions in Europe and are important as a quarry animal to the Inuit. Most commercial venison in the United States is imported from New Zealand.

Deer were originally brought to New Zealand by European settlers, and the deer population rose rapidly. This caused great environmental damage and was controlled by hunting and poisoning until the concept of deer farming developed in the 1960s. Deer farming has advanced into a significant economic activity in New Zealand with more than 3,000 farms running over 1 million deer in total. Deer products are exported to over 50 countries around the world, with New Zealand becoming well recognised as a source of quality venison and co-products.

White-tailed deer hunted in Accomack, Virginia

Automobile collisions with deer can impose a significant cost on the economy. In the U.S., about 1.5 million deer-vehicle collisions occur each year, according to the National Highway Traffic Safety Administration. Those accidents cause about 150 human deaths and $1.1 billion in property damage annually.[36] In Scotland, several roads including the A82, the A87 and the A835 have had significant enough problems with deer vehicle collisions (DVCs) that sets of vehicle activated automatic warning signs have been installed along these roads.[37]

The sight of deer standing motionless, caught in headlights gives rise to the phrase "deer in the headlights".

In some areas of the UK, deer (especially fallow deer due to their gregarious behaviour), have been implicated as a possible reservoir for transmission of bovine tuberculosis,[38][39] a disease which in the UK in 2005 cost £90 million in attempts to eradicate.[40] In New Zealand, deer are thought to be important as vectors picking up M. bovis in areas where brushtail possums, (Trichosurus vulpecula), are infected, and transferring it to previously uninfected possums when their carcasses are scavenged elsewhere.[41] The white-tailed deer, (Odocoileus virginianus), has been confirmed as the sole maintenance host in the Michigan outbreak of bovine tuberculosis which remains a significant barrier to the US nationwide eradication of the disease in livestock. In 2008, 733,998 licensed deer hunters harvested approximately 489,922 white-tailed deer in attempts to control the deer population and disease spread. These hunters purchased more than 1.5 million deer harvest tags. The economic value of deer hunting to Michigan's economy is substantial. For example, in 2006, hunters spent US$507 million hunting white-tailed deer in Michigan.[42]

Deer hunting is a popular activity in the U.S. and generates revenue for states and the federal government from the sales of licenses, permits and tags. The 2006 survey by the U.S. Fish and Wildlife Service estimates that license sales generate approximately $700 million annually. This revenue generally goes to support conservation efforts in the states where the licenses are purchased. Overall, the U.S. Fish and Wildlife Service estimates that big game hunting for deer and elk generates approximately $11.8 billion annually in hunting-related travel, equipment and related expenditures.[43]